Lec. 5 WATER Uptake 4-1. Driving forces for water flow: 1. How does water move into the root? 3 main water Apoplastic pathway potential gradients Symplastic & transmembrane pathway Water enters the cytoplasm at the endodermis 2. How is water transported from roots to the leaves? Pressure-driven bulk flow. Evidence for a negative pressure in xylem. Negative pressure is generated by transpiration. Water moves up a plant by the Cohesion-Tension Theory

4-4. A. Water uptake pathways across the root Endodermis: a layer of cells that prevents ion and water loss. Because of the Casparian strip, a suberin (waxy) barrier, water enters the cytoplasm at the endodermis (PM contains water channels or aquaporins. Water channel activity may be regulated.) Water then leaves the endodermis through the plasma membrane and moves to the xylem using both the apoplast and symplast pathways.

Root Pressure: positive pressure in the root xylem due to ion accumulation

4-10. Driving force for water B. Water is conducted to leaves via the xylem loss: by pressure-driven bulk flow. water vapor conc. difference 4-7. Two types of xylem How much driving force is needed?

1 4-9. Tension or negative pressure originates in leaves

4-9. Negative pressure of water on curved surfaces depends on radius 4-7. Tension in xylem can cause cavitation. Ψp = -2T/r

Other theories: Xylem pressure not – enough, may be a technical limitation. Estimate: Cohesion-Tension Need 3 Mpa to theory still stands lift water up 100 Steudle 2001 m in xylem of ARPB 40 um radius.

How do we know there is a negative pressure in xylem Pull: Water is under negative pressure. or in cells? Push: Water is under positive pressure Expt. Evidence 1. Pressure chamber 2. Pressure probe (single cell) (see Web: www.plantphys.net)

How is the negative pressure generated? a) Transpiration: loss of water vapor through stomatal pores b) Growing cells that take up water Water moves up by cohesion-tension theory

2 Review: Waters enters the root through the apoplast, symplast and -95 Mpa transmembrane pathways. -0.8 Water moves from roots to leaves by pressure-driven bulk flow in tube-like structures- xylem. -0.8 Negative pressure is generated by transpiration. Transpiration is the loss of water through stomatal pores. -0.6 Negative pressure is also formed by growing cells that take up -0.3 water.

Negative pressure is also generated by How is water potential measured? growing cells that take up water 1. Osmotic or solute potential of tissues. Tissue weight as a function of solute concentration 2. Xylem pressure potential When humidity is high Pressure chamber When stomatal aperture is Pressure probe closed 3. Tissue water potential Thermocouple psychrometer

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